24
NEW AND LITTLE KNOWN MARINE NEMATODES FROM A SCOTTISH SANDY BEACH by Richard M. Warwick (*) Dove Marine Laboratory, Cullercoats, Northumberland, England and Howard M. Platt (**) Department of Zoology, Queen's University, Belfast, N. Ireland. Résumé Treize espèces de Nematodes marins libres de la plage de sable de Firemore Bay, Wester-Ross, Ecosse, sont décrites. Parmi ces espèces, nous proposons un nouveau genre, Gairleanema, avec l’espèce-type G. anagremilae gen. et sp. nov. Onze autres sont de nouvelles espèces attribuables aux genres existants : Odon- tophora exharena sp. nov., Theristus (Daptonema ) gelana sp. nov., Cobbia Cale donia sp. nov., Gonionchus inaequalis sp. nov., Xgala smo sp. nov., Siphonolaimus ewensis sp. nov., Metachromadora (Bradglaimus) scotlandica sp. nov., Microlaimus acinaces sp. nov., Dasgnemella albaensis sp. nov., Rhabdodemania imer sp. nov. et Trileptium parisetum sp. nov. Une espèce ( Trefusia zostericola Allgén, 1933), jusqu’ici peu connue est redécrite. INTRODUCTION In July 1971, a study of the meiofauna of the sandy beach at Firemore Bay (Gaineahm Smo), Loch Ewe, Wester-Ross, Scotland, was undertaken. The location, physical characteristics and macro- faunal population of this beach are described by McIntyre and Elef- theriou (1968). Several workers will collaborate in the subsequent publication of the ecological results but we here take the opportunity of describing the new, or in one case poorly known, nematode species which were found during the investigation. Descriptions have been made from glycerine mounts and the type material is deposited at the British Museum (Natural History). Curved structures such as spicules have been measured as the chord and not the curve. Following convention, we have given both Filip- (*) Present address: Institute for Marine Environmental Research, 13-14 St James Terrace, Citadel Road, Plymouth, PLI 3AX, Devon, England. (**) Present address : British Antarctic Survey, Monks Wood Experimental Station, Abbots Ripton, Huntingdon, England. C ahiers db biologie marine Tome XIV - 1973 - pp. 135-158.

NEW AND LITTLE KNOWN MARINE NEMATODES - VLIZ · NEW AND LITTLE KNOWN MARINE NEMATODES FROM A SCOTTISH SANDY BEACH. by. Richard M. Warwick (*) Dove Marine Laboratory, Cullercoats,

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NEW A N D LITTLE KNOW N MARINE NEMATODES

FROM A SCOTTISH SANDY BEACH

by

Richard M. Warwick (*)Dove M arine L aboratory, Cullercoats, N orthum berland, England

and

Howard M. Platt (**)D epartm en t of Zoology, Q ueen 's University, Belfast, N. Ireland.

Résumé

Treize espèces de Nematodes marins libres de la plage de sable de Firemore Bay, W ester-Ross, Ecosse, sont décrites. Parmi ces espèces, nous proposons un nouveau genre, Gairleanema, avec l ’espèce-type G. anagremilae gen. et sp. nov. Onze autres sont de nouvelles espèces attribuables aux genres existants : Odon- tophora exharena sp. nov., Theristus (Daptonema) gelana sp. nov., Cobbia Cale­donia sp. nov., Gonionchus inaequalis sp. nov., Xgala smo sp. nov., Siphonolaimus ewensis sp. nov., Metachromadora (Bradglaimus) scotlandica sp. nov., Microlaimus acinaces sp. nov., Dasgnemella albaensis sp. nov., Rhabdodemania imer sp. nov. et Trileptium parisetum sp. nov. Une espèce (Trefusia zostericola Allgén, 1933), jusqu’ici peu connue est redécrite.

INTRODUCTION

In Ju ly 1971, a study of the meiofauna of the sandy beach at F irem ore Bay (Gaineahm Smo), Loch Ewe, W ester-Ross, Scotland, was undertaken. The location, physical characteristics and macro- faunal population of this beach are described by McIntyre and Elef- theriou (1968). Several workers will collaborate in the subsequent publication of the ecological results but we here take the opportunity of describing the new, or in one case poorly known, nematode species which were found during the investigation.

Descriptions have been made from glycerine m ounts and the type m aterial is deposited at the British Museum (Natural H istory). Curved structures such as spicules have been m easured as the chord and not the curve. Following convention, we have given both Filip-

(*) Present address: Institute for Marine Environmental R esearch , 13-14 St James Terrace, Citadel Road, Plymouth, PLI 3AX, Devon, England.

(**) Present address : British Antarctic Survey, Monks Wood Experimental Station, Abbots Ripton, Huntingdon, England.C a h i e r s d b b i o l o g i e m a r i n e Tome XIV - 1973 - pp. 135-158.

136 R.M. WARWICK AND H.M. P L A T T

jev’s form ula and de Man’s ratios as a prelude to each description. The application of both these conventions is outlined by Vitiello (1969).

Family AXONOLAIMIDAE

ODONTOPHORA EXHARENA sp. nov. (Fig. 1)

Material studiedSyntypes : 1 $ and 1 9, B.M. (N.H.) Reg. Nos. 1972:331.

Fio. 1Odonthophora exharena sp. nov.

A: male head; B : fem ale head; C: male ta il.

Body formulae and ratios2,167

¡$ :

9 :

— 144 M15 23 23 24

— 162 1,360 2,18015 23 28 18

2,280pm; a = 99.1; b = 15.8; c = 20.2;

Spicule = 30 (j.m

2,300tmi; a = 82.1; b = 14.2; c = 19.2;

V p. 100 = 59.1

MARINE NEMATODES 137

Description

Cuticle w ith fine transverse striation. Six labial papillae small and indistinct. Four cephalic setae 19-21 pm. Subcephalic setae in two whorls. Four setae 9-11 pm long in submedian positions about half way down the length of the amphids. Second whorl of four 11-13 pm sublateral setae and four 2.5 pm submedian setae ju st pos­terior to amphids. Buccal cavity typical of the genus. Odontia 7 pm long; posterior portion of buccal cavity conical, 13 pm long. Amphids in form of a large open loop 16 pm long and 7.5 pm wide. Oesophagus w ith no posterior bulb, nerve ring 102-104 pm from anterior. Somatic setae up to 13 pm long rather sparsely scattered down length of oesophagus, middle region of body virtually naked. Tail conical in both sexes, 4.7 cloacal diameters long in male and 6.7 anal diameters in female.

Male: Spicules paired, uniformly arcuate, cephalated proximally and pointed distally. Gubernaculum with paired dorsal apophyses, 19 pm long. Small papilliform pre-cloacal supplements indistinct, but there appear to be only four.

Fem ale: Ovaries paired, symmetrical, opposed and not reflexed.

Discussion

Odontophora exharena sp. nov. is distinguished from other mem­bers of the genus by the combination of very large amphids, long cephalic setae and the arrangem ent of subcephalic setae.

Fam ily MONHYSTERIDAE

THERISTUS (DAPTONEMA) GELANA sp. nov.(Fig. 2)

Material studied

Syntypes: 3 4 4, B.M. (N.H.) Reg. Nos. 1972: 332-333.

Body formulae and ratios

1,095pm; a = 28.8; b = 4.3; c = 8.2;— 253 M 96023 38 38 30

— 260 M 95024 39 39 30

— 241 M 80522 37 37 30

Spicule = 45 pm

1,080pm; a = 27.7; b = 4.2; c = 8.2;

Spicule = 46 pm

915pm; a = 24.7; b = 3.8; c = 8.5;

Spicule = 44 pm

138 R,M, WARWICK AND HM, P L A T T

Description

Cuticle distinctly transversely striated. Six internally striated lips and six 2 $xm long labial setae. Six 16-17.5 ji.m cephalic setae, four 14 um submedian cephalic setae and occasionally a th ird seta of interm ediate length (16jj.ni) may be distinguished in the submedian groups. All three setae in these submedian groups are usually diffi­cult to distinguish as they are regularly closely applied to one another. There are eight files of long fine cervical and somatic setae ; 35-65 pm cervically, 50-65 jun in the middle regions and 5-35 jim caudally. Paired term inal setae 25-35 jj.ni.

F ig . 2Theristus (Daptonema ) gelana sp. nov.

A: male head; B: amphid of another m ale; C: lateral view of spicules and gubernaculum; D: male tail.

Buccal cavity conical, unarm ed. Amphid situated 25 jim from the anterior, 13-14 jim wide, with a corresponding body diam eter of 29-30 %im. In most specimens the structu re of the am phid is very indistinct, but in one specimen was seen as a relatively distinct spiral structure of 2.5 turns (Fig. 2 B). Oesophagus lacks a posterior bulb. Tail conical in anterior part, then cylindrical; 3.7-4.5 cloacal body diam eters long.

Male: Spicules paired, equal, w ith a distal hook; 1.5 cloacal body diameters long. The gubernaculum is an inter-spicular plate bearing complicated pincer-like lateral pieces.

MARINE NEMATODES 139

Discussion

W e have experienced some difficulty in the generic placing of this species. If we use the criteria of W ieser (1956), it should be placed w ith Paramonhystera or Promonhystera because of the indis­tinct thin-walled amphids. However, in both these genera the males have long slender spicules. Furtherm ore, all species of Paramon­hystera w ith long somatic setae have these confined to a single circle ju s t behind the amphids (subgenus Leptogastrelia Cobb, 1920), and all species of Promonhystera have very long labial setae. Hopper (1968) transfers Promonhystera albigensis Riemann, 1966 to the subgenus Daptonema of the genus Theristus on the basis of the structu re of the male genital apparatus alone. He argues th a t this species has a gubernaculum similar to the group centred around Theristus (D aptonem a) beutschlii, for which he provides a key. The present species also has a gubernaculum sim ilar to other members of this group, and we therefore provisionally place it w ith Daptonema. The fact th a t other short spiculed species are found in the group adds weight to this argum ent. The only other species in the group with somatic setae longer than the corresponding body diam eter is T. (D.) albigensis, which is immediately distinguished from T. (D.) gelana by its long slender spicules.

COBBIA

Holotype: 1 S, B.M. (N.H.)

F ig . 3C obbia Caledonia sp. nov.

A: male head; B: male tail; C: lateral view of spicules.

Body formulae and ratios216 M 925

3 : -J4---- 20 22-----20 I .°00(xm approx.?; a = 45; b = 4.6; c = 14?;Spicules = 25 tun

CALEDONIA sp. nov. (Fig. 3)

arial studied

, No. 1972:334.

140 R.M. WARWICK AND H.M. P L A T T

Description

Cuticle distinctly striated. Labial setae 2.5 ¡mi. Six longer cephalic setae 15 ¡mi; four shorter submedian cephalic setae 11 ¡mi. Somatic setae relatively short (5-10 ¡ma) and sparse. Buccal cavity contains three lightly cuticularised teeth, the dorsal one extending further anteriorly than the two subventral ones. Amphid 34 ¡mi from anterior, 11 ¡mi wide and 12 ¡mi long, corresponding body diam eter 18 ¡mi. Tail conical but most probably broken, the long flagellum found in m any similar species having been lost.

Male: Spicules paired, equal, cephalate proximally. Guberna- culum a very lightly cuticularised and indistinct structu re lying parallel to the distal portion of the spicules.

Discussion

A key to the genus is given by W ieser (1959). This species belongs to the group in which the dorsal tooth is longer than the subventral. It is distinguished from other m em bers of the group m ainly by the size and position of the am phid and also by the combination of the lengths of the labial setae, cephalic setae and spicules as outlined in the following table :

. C. Caledonia C. dentata C. urinator C. trefusiaformtsC h a ra c te r sp. nov Gerlach, 1953 W ieser, 1959 de Man, 1907

Body le n g t h approx. 1000 Mm 1000 Mm 1590 Mm 2000 AtmLabial setae length ............... 2.5 Atm 6 Atm 6 Atm 4 Atm

Cephalic setae length^ & ) . . 15 / tm (1.1) 22-26 Atm (1.5) 25 Atm (1.3) 25.5 Atm (1.1)Amphid from the anterior2. 34 Atm (2.4) 22-26 Atm (1.5) 25 Atm (1.3) — (1.5)Width of amphid3................... 11 Atm (61) 10 Atm (50) 8.5 Atm (33) 8 .7 /tm (33)Spicule length4 .......................... 25 Atm (1.3) 20 Atm (1.0) 36 / t m (1.1) 40 Atm (1.0)

1 . . . . length as fraction of head diameter in parenthesis2 . . . . length as fraction of head diameter in parenthesis3 . . . . w idth as percentage body diameter in parenthesis4 . . . . length as a fraction of the cloacal diameter in parenthesis.

G O N IO N C H U S INAEQUALIS sp. nov. (Fig. 4)

Material studied

Syntypes: 1 $ and 1 9 , B.M. (N.H.) Reg. Nos. 1972: 335.

Body formulae and ratios

1,375 ¡im; a =

Spicule = 34 + 46 ¡un

1,680 um; a = -

V p. 100 = 74.

* 377 M 1,127 1 o u o a . % %.a : —----- —— —-------— 1,375 ¡im; a = 39.3; b = 3.6; c = 5.5;

Q — 477 M 1,247 1,360 t _on , Q , , „S: 22' 37 42-------39------30 1’68° !*D1: * = 4° ° : b = 3'5> C = 5'3 ’

MARINE NEMATODES 141

Description

Cuticle transversely striated ; no lateral differentiation observed. Six well-developed lips, each with a characteristic triangular flange. Labial setae 4-4.5 pm, seemingly situated on conical extensions of the labial cuticle. Six 22-26 pm cephalic setae plus ten other 7-11 pm setae (four subm edian pairs and two lateral). Four short 5 pm sub- cephalic setae are situated a short distance behind the cephalic setae. Eight files of cervical setae begin level with the amphids and are most

F ig. 4Gonionchus inaequalis sp. nov.

A: male head; B: lateral view of spicules and gubernaculum; C: male tail.

num erous in the region immediately behind the amphid ; length 7-140 pm, i.e., up to 400 p. 100 of the corresponding body diameter. Somatic setae as far as the anus or cloaca are spaced 60-120 pm apart longitudinally and are 250-300 p. 100 of the corresponding body diam eter in length. Caudal setae only sparse and short; long ter­m inal setae apparently absent.

Around the widest part of the buccal cavity is a narrow cuticu- larised band, which has six short double pegs extending anteriorly into the bases of the lips. Two subventral cuticularised flanges are present in the buccal cavity. The circular amphid is ra ther indis-

142 R.M. WARWICK AND H.M. P L A T T

linci, in the male 7 ¡im (28 p. 100 of the corresponding body diam eter) wide and 22 ^m from the anterior ; in the female 9 ^m (33 p. 100 of the corresponding body diameter) wide and 21 *im from the anterior. The long tapering tail is 8.0 cloacal body diam eters long in the male and 10.5 anal body diameters in the female.

Male: Spicules paired, unequal ; the right 1.5 cloacal body dia­meters long and the left 1.1 cloacal body diam eters. Both are cepha- late proximally. Paired dorsal gubernacular apophyses 11 ^m long.

Female: Ovary single, anterior. Eggs 90 X 35 ¡¿m.

Discussion

Cobb (1920) describes Gonionchus villosus, which has two sub­median plate-like onchia, “term inal flaps... hinged to the lips” and long slender somatic setation. Although the illustration he gives is rather poor, the description is sufficient for the present species to be placed in this genus. G. inaequalis sp. nov. is distinguished from the type by the possession of unequal spicules and gubernacular apophyses.

XYALA SMO sp. nov.(Fig. 5)

Material studied

Syntypes: 2 S $ and 2 9 5, B.M. (N.H.) Reg. Nos. 1972: 336-337. Paratype: 1 &, personal collection of R.M.W.

Body formulae and ratios

1,356{xm; a = 43.7; b = 4.4; c = 7.5;— 311 M 1,17619 29 31 25

— 306 M 1,01319 29 30 23

— 360 993 1,20722 30 31 24

— 316 900 1,09721 29 31 23

Spicule = 30 [xm

1,319 urn; a = 44.0; b = 4.3; c = 7.5;

Spicule = 29 ijim

1,407 %im; a = 47.4; b = 4.1 ; c = 5.6;

V p. 100 = 67.5

1,310pm; a = 42.3; b = 4.1; c = 6.1;

V p. 100 = 68.5

Description

Cuticle transversly striated and also m arked w ith longitudinal files of cuticular blocks. Anteriorly these blocks carry characteristic dumbell shaped excrescences the height of which gradually reduces posteriorly until approximately half way along the length of the oesophagus they are no longer visible. The num ber of longitudinal

MARINE NEMATODES 143

files in the head region is 32, reducing to about 24 in the middle region of the body and 20 in the cloacal or anal region.

The m outh is surrounded by six characteristic high and rather hyaline lips which bear short 3-4 pun labial setae. At the anterior end of the somewhat conical buccal cavity six V-shaped cuticular ridges extend forw ards into the lips. There are 6 pairs of unequal cephalic setae, 24-28 pun and 18-21 pun in length. The tips of these setae have a withered appearance. There are four very short sub-

Fig. 5Xyala smo sp. nov.

A: male head; B: cuticular pattern at base of oesophagus; C: lateral view of spicules and gubernaculum; D: male tail; E: female tail.

median setae on the head ju st posterior to the cephalic setae. There are a few scattered setae in the cervical region, up to 29 pun in length. The am phids are th in walled and somewhat ovoid, 7 pun wide in the male and 6 pun in the female, and are situated 25-31 ¡un from the anterior. The corresponding body diameter is 23 ¿im in the male and 26 ¡pun in the female. The oesophagus is cylindrical throughout its length, w ith no posterior bulb. The tail is of a different shape in the two sexes. In the male it is 7.2-7.6 cloacal body diameters long, with the anterior two thirds tapering and the posterior th ird cylin-

3

144 R.M. WARWICK AND H.M. P L A T T

drical. In the female it is 9.3-11.0 anal body diam eters long and more evenly tapered throughout its length. In addition the male tail bears 2 subventral rows of short (4 pm) setae.

Male: The spicules are paired, equal, 1.2-1.3 cloacal body dia­m eters long. They are sharply bent in the middle and cephalate proximally. The gubernaculum is in the form of an inter-spicular plate with a weakly cuticularised dorsal apophysis.

Female: There is a single anterior ovary, but the details of its structure are not clear.

Discussion

Currently two other species are referred to this genus ; Xyala striata Cobb, 1920 and X. exigua W ieser, 1956. X. smo is distinct from these on many points, among which m ay be noted the head structure, relative lengths of labial and cephalic setae, cuticular pattern and shape of the tail.

Family SIPHONOLAIMIDAE

SIPHONOLAIMUS EWENSIS sp. nov.(Fig. 6)

Material studied

Syntypes: 2 S S, B.M. (N.H.) Reg. Nos. 1972:338.Paratypes : 5 Juveniles, personal collection of R.M.W.

Body formulae and ratios

ê : ~ 2H ^ 3’7^ 3,910 tun; a = 106; b = 14.5; c = 32.1;17 33 37 39

— 260 M 4,23717 35 37 39

Spicule = 63 pm

4,370pm; a = 1 1 7 ; b = 16.8; c = 32.9;

Spicule = 56 pm

Description

Cuticle transversely striated. Ten cephalic setae; submedian pairs 18-22 pm + 5 pm, lateral setae 9 pm. Circle of 6 subcephalicsetae level w ith anterior border of amphids, 17 pm long. Amphids thick walled, circular, 17-18 pm diam eter (62-71 p. 100 of corres­ponding body diam eter), attachm ent of am phidial nerve prom inent.Buccal cavity with axial spear 56-63 pm long. Posterior half of oeso­phagus dilated. Nerve ring 42-46 p. 100 of oesophagus length from anterior. Intestine full of opague black granules. Setae sparse on mid-body region, becoming more num erous on the male tail. Tail conical, 3.1-3.4 cloacal body diam eters long.

MARINE NEMATODES 145

Male: Spicules 1 .4-1 .6 cloacal body diameters long, arcuate, cephalate proxim ally and with weakly cuticularised ventral alae. The dorso-caudal apophysis of the gubernaculum is 27-30 pm long. On either side of the gubernaculum there is a well cuticularised lateral swelling w ith a cup-shaped distal end. There are six promi­nent setose pre-cloacal supplements.

Fig. 6Siphonolaimus ewensis sp. nov.

A: male head; B: male ta il; C, D: spicules and gubernaculum of two males.

Discussion

Siphonolaimus ewensis sp. nov. is closest to S. cobbi Riemann, 1966. This latter species, however, has a very differently shaped gubernaculum lacking cup-shaped lateral pieces. Several other fea­tures which m ay be used to separate the two species are set out below.

S. cobbi S. ewensisspicule 38.5 p.m spicule 56-63 pm

tail 2.3 anal body diameters tail 3.1-3.4 anal body diameters c = 60 c = 32.1-32.9

spear length 36 pun spear length 56-62 pm3 supplem ents 6 supplements

146 R.M. WARWICK AND HM. P L A T T

F am ily SPIRINIDAE

METACHROMADORA (BRADYLAIMUS) SCOTLANDICA sp. nov.(Fig. 7)

Material studiedSyntypes : 3 S B.M. (N.H.) Reg. Nos. 1972: 339-340.

F ig . 7Metachromadora (Bradyla im us)

scotlandica sp. nov.A: male head; B: oesophageal bulb; C: lateral view of am- phid; D: male ta il; E: lateral view of spicules and guberna- culum.

— 178 M 1,47324 32 40 35

— 175 M 1,07523 36 21 32

— 200 M 1,18323 36 33 34

Body formulae and ratios

1,570pm; a <=39.3; b = 8.8; c = 16.2;

Spicule = 31.5 pm

1,140pm; a = 36.8; b = 6.5; c = 17.5;

Spicule = 34.5 pm

1,270 pm; a = 38.5; b = 6.4; c = 14.7;

Spicule = 36 pm

DescriptionCuticle w ith fine transverse striation. Six labial setae 3-4 pm

long. Cephalic setae comprise an anterior circle of four and a slightly

MARINE NEMATODES 147

more posterior circle of six 12.5-14 pm setae . There are a few short scattered setae (4-5 pm) level with the amphids, and more poste­riorly two circles each of 6-7 setae, about equal to the cephalic setae in length. Setae are sparse and short on the rem ainder of the body. The amphids form a single open loop 8.5-9 pm wide, the corresponding body diam eter being 27-29 pm. The buccal cavity contains a large, hollow, strongly cuticularised dorsal tooth, opposed by two very small subventral projections. The oesophagus has an elongate posterior bulb 59-68 pm long and 27-31 pm wide. The bulb has a thick cuticular lining which is divided in the middle by a constriction. The tail is conical, 2.0-2.8 cloacal body diameters long. U nstriated tail tip 17 pm long.

Male: Spicules paired, equal, arcuate, slightly cephalate proxi- mally and w ith ventral alae. They are about 1 cloacal body diameter in length. Gubernaculum 19-23 pm, crescentic. There are 11-13 pre- cloacal supplem ents each consisting of a conical elevation of the cuticle w ith a narrow duct leading to the interior. The supplements are less outstanding when the posterior part of the body is not ventrally bent. The anteriorm ost supplement is 127-206 ¿on in front of the cloaca. The tail bears a ventral supplement, 21-37 pm behind the cloaca, which is of the same structure as the pre-cloacal supple­m ents. Two subventral files of short setae are present on the posterior p art of the body, commencing at about the level of the anterior supplem ent.

Discussion

W ieser & Hopper 1967 provide a useful key to the subgenera and species of Metachromadora. The present species is closest to M. ger- lachi (Gerlach, 1955) W ieser & Hopper, 1967. However, in M. gerlachi there is no tail supplem ent and the spicules are square ended and not roundly cephalate. Furtherm ore, the two species have a different arrangem ent of cephalic setae, and the oesophageal bulb in M. ger­lachi is 82 pm long (as opposed to 59-68 pm in M. scotlandica) w ith its cuticular lining divided into two completely separate sections.

Fam ily MICROLAIMIDAB

MICROLAIMUS ACINACES sp. nov.(Fig. 8)

Material studied

Syntypes : 3 3 4, B.M. (N.H.) Reg. Nos. 1972:341.

Body formulae and ratios

13 % 30----- 1,150t<.m; a = 38.5; b = 8.3; c = 11.9;Spicule = 27 pm

148 R.M. WARWICK AND HM. P L A T T

$ : ~~-----------^ ----- 3̂ 1,010^m; a = 34.8; b = 9.3; c = 11.9;13 28 29 22

Spicule $= 28 ixm

3: ~ ^ 7 " 2 9 1,0fi 1.095(xm; a = 37.8; b = 8.2; c = 15.0;Spicule *= 28 ¡im

F ig . 8Microlaimus acinaces sp. nov.

A: m ale head; B: lateral view of spicules and gubernaculum ; C: m ale tail.

Description

Cuticular striations commence ju s t anterior to the amphids. Head cuticle is restricted posterior to the cephalic setae. Six 2-2.5 y.m and four 11-12 yum cephalic setae in two circles. Four 7-8 yun cervical setae behind amphids. Caudal setae, in male, in two subventral files; 7-8 jun long. Buccal cavity well developed w ith a large dorsal and two smaller subventral teeth. Peribuccal tissue well developed. Amphids situated 10-11 pm from anterior, 8 y.m wide; corresponding body diam eter 13-14 yon. Oval posterior oesophageal bulb about 19 X 25y.m. Tail conical, 3.0-4.0 cloacal body diam eters long.

Male: Spicules paired, equal and sickle-shaped; 1.1-1.3 cloacal body diam eters long. Paired gubernaculum 21-22 yun long, proxim ally spoon-shaped. Four small preanal supplem ents present. There are two files of nine subventral pre-cloacal cuticularised ducts which em anate from subcuticular glands.

Discussion

This species belongs to Gerlach’s (1950) group A on the basis of the well developed buccal cavity. It is probably closest to M. paraborealis Allgén, 1940 sensu Gerlach, 1950, on the basis of the size and position of the amphids and presence of postanal setae. The two species differ, however, in the length of the cephalic and postanal setae, the length of the gubernaculum and the form of the spicules. M. acinaces is further characterised by the presence of pre- cloacal supplements and subventral glands.

MARINE NEMATODES 149

Family DASYNEM ELLIDAE

DASYNEMELLA ALBAENSIS sp. nov.(Fig. 9)

Material studiedSyn types : 1 d and 2 9 $ , B.M. (N.H.) Reg. Nos. 1972: 342-343.

ç

F ig . 9Dasynemella albaensis sp. nov.

A: male head; B: fem ale head; C: female tail; D: lateral view of spicules and gubernaculum.

d :

9 :

9 :

— 204 M 1,48313 19 22 20

— 179 765 1,33613 21 27 17

— 203 780 1,37814 21 29 21

Body formulae and ratios

1,610am; a = 7 3 .2 ; b = 7.9; c = 1 2 .7 ;

Spicule = 24 [im

1,430tun: a = 53.0; b = 8.0; c = 11.5;

V p. 100=53.5

1,500 (im; a = 51.7; b = 7.4; c = 1 2 .3 ;

V p. 100=52.0

150 RM, WARWICK AND H.M, PL A T T

Description

Cuticle m arked by coarse transverse striations and bears longi­tudinal files of obliquely orientated cuticular flanges. There are 8 files on the anterior and posterior parts of the body, 10 on the middle region and 6 near the tail tip. No setae on the general body surface. Cuticle in the head region m arkedly thickened form ing a cephalic capsule 27-32.5 {¿in long and 17-18 ¡jun wide at its base. Capsule unstriated but bears eight cuticular ribs which extend for­wards from its base. The capsule is slightly constricted posterior to the cephalic setae, and is pierced by median pores at the level of the amphids. An anterior circle of six 7-10 ^m cephalic setae and more posteriorly four 10-15 ^m cephalic setae. The head diam eters given in the above formulae are m easured at the level of these posterior setae. Buccal cavity absent. Amphids form a closed loop and are oval in outline (5 jjjk wide) in the male, but circular (6-6.5 ¡mi wide) in the female. Amphids 15-22 ^m from anterior. Oesophagus narrow , cylindrical, no posterior bulb. Tail conical, 5.S-7.3 anal or cloacal body diam eters in length. Unstriated tip 22.5-31 jmi long, w ith six cuticular ribs extending two thirds of the way down its length.

Male : Spicules 1.2 cloacal diam eters long, ra ther weakly cuticu- larised especially at the proximal end. Gubernaculum a weakly cuti- cularised plate 15 ^m long.

Fem ale: Because of the thick body cuticle, details of the female reproductive system are not clearly visible.

Discussion

This species is distinguished by the position and form of the amphids, the lengths of the cephalic setae and the cuticular pattern . Riemann (1966) describes a species of Dasynemella which is alm ost certainly identical with D, albaensis. However, he had no males and was unable to determine the form of the am phids, and for these reasons did not feel justified in establishing a new species.

Family OXYSTOMINIDAE

TREFUSIA ZOSTERICOLA Allgén, 1933 (Fig. 10)

Material studied1 a.

Body formulae and ratios99(1 9 991

23— 30 30 ' s o 3’550*m; a c l l 8 0 ; b = = 1 0 8 ’ c = 16-2;Spicule = 55 |xm

MARINE NEMATODES 151

Description

Six 14 (im jointed cephalic setae. Subcephalic setae 9 y.m, situated posterior to amphid (36 tun from an terio r). Two lateral setae 34 and 52 (im behind amphids. Amphid 24 am from anterior, maximum width 6 p.m (corresponding body diameter 24 p.m). Tail 7.3 cloacal diam eters long.

F ig. 10Trefusia zostericola Allgén, 1933

A: male head; B: m ale ta il; C: lateral view of spicules and gubernaculum.

Male: Spicules paired, equal, slightly cephalate proxim ally; 1.8 cloacal diam eters long. Gubernaculum a slender rod with two well developed lateral pieces. Pre-cloacal supplements present.

Discussion

An excellent discussion and key to this genus is given by Riemann (1966a). This species is characterised by the elongate spicules, the tail length and the position of the subcephalic setae. Only Allgén

152 R.M. WARWICK AND H.M. PL A T T

has described and recorded T. zostericola previously; his figures are rather poor, but sufficient to allow the present specimen to be equated with it.

Family ENOPLIDAE

GAIRLEANEMA ANAGREMILAE gen. et sp. nov.(Fig. 11)

Material studied

Syntypes 2 « Ä, B.M. (N.H.) Reg. Nos. 1972: 344-345.

F ig. 11Gairleanema anagremilae gen. et sp. nov.

A: male head; B: lateral view of spicules and gubernaculum ; C: male tail.

Body formulae and ratios

OT “ 1,838 1 . 9 8 5 ^ ; a = 45.1; b - 5 . 7 ; c = 10.9:

Spicule s= 52 %im

41 1,738 1 .8 8 0 ,m ; a = 46.1; b = 5.7; c = 1 2 . 8 :

Spicule = 53 y.m

MARINE NEMATODES 153

Description

Cuticle w ith ra ther indistinct transverse striation. Mouth sur­rounded by three high lips each w ith a pair of 28-30 puin labial setae. The labial setae have bulbous swelling about half way along their length. Ten cephalic setae; six longer ones 68-69 pm w ith a thin section near the tip ; four shorter 7 y.m. There are a few short (up to4.5 ixm) setae scattered on the anterior cervical region. Caudal setae and two subventral 17 (im term inal setae present, bu t other somatic setation apparently lacking. Cephalic capsule weakly developed with its posterior border indistinct. Buccal cavity w ith a pointed heavily cuticularised dorsal onchium pierced by a duct which leads into the oesophageal m usculature. Two subventral teeth less strongly cuticu­larised, w ith rounded tips. Amphids situated ju st behind lateral cephalic setae, indistinct but with the point of junction of the amphi- dial nerve prom inent. There is a small (7-8 pm diam eter) circular area of unstriated cuticle situated laterally ju st behind the cephalic capsule. Oesophagus cylindrical, 22-25 jxm wide at its base. Nerve ring 144-150 pun from anterior. Tail 3.0-4.8 cloacal diameters long, posterior half cylindrical.

Male: Spicules 1.4 cloacal diameters long, equal, L-shaped, cepha- late proxim ally and w ith broad ventral alae. Gubernaculum paired, 21 pun long, w ith hooked dorsal apophyses. There is a ventral pre- cloacal seta 11.5-13 pun long, a small ventral papilla 12.5-13 ¡un in front of the cloaca and two large rounded mammilate protuberances in front of this. The posterior protuberance is 32 pun in front of the cloaca and the anterior 43-48 pun in front of the cloaca. The nipples are pierced by fine pores.

Discussion

Gairleanema gen. nov. is probably closest to Saveljevia Filipjev, 1927, which has well developed onchia and vestigeal or reduced mandibles. This la tte r genus is poorly known, since all its species (S. curvidens Filipjev, 1927; S. kolaensis Filipjev, 1927; S. xiphonchus Filipjev, 1927; S. bastata Wieser, 1953; S. cornuta Gerlach, 1956) are described from females or juveniles only. However, each of these species has cephalic setae which are considerably shorter than one head diam eter, simple non-bulbous labial setae, and most im portantly the dorsal onchium is small or reduced. None of these features perta in in Gairleanema, in which the dorsal onchium is large and more heavily cuticularised than the subventral ones. Gairleanema is further distinguished from other genera of the Enoplidae by the unique struc­tu re of its pre-cloacal supplements, which in other genera are almost always single, tubu lar and cuticularised. The relationship between Gairleanema and Saveljevia cannot be elucidated until males of the la tte r genus are found. The generic diagnosis of Gairleanema is as follows :

Enoplidae; m andibles absent; dorsal onchium not reduced in size; labial setae w ith bulbous swellings ; pre-cloacal supplements m am m illiform .

154 R.M. WARWICK AND H.M. P L A T T

RHABDODEMANIA IMER sp. nov.(Fig. 12)

Material studied

Holotype : 1 ê , B.M. (N.H.) Reg. No. 1972: 346 plus 1 9 in very poor condition and not used in description.

F ig . 12Rhabdodemia imer sp. nov.

A: male ta il; B : male head; C: lateral view of spicules and gubernaculum.

Body formulae and ratios

2,960 ¡un; a = 12

Spicule = 26 ¡¿in

cf : IQ ^ 2’822l 2,960 tun; a = 123.3; b = 8.0; c = 21.1;

MARINE NEMATODES 155

Description

Cuticle smooth. Six swollen rounded lips each w ith a small labial papilla. Cephalic setae in two separate circles, anterior four 4 pm, posterior six 16.5 ^m. Head diameter at level of longer setae12.5 %im. Buccal cavity conical, appears to contain three weakly developed teeth a t its anterior end. Four 8 $im cervical setae some distance posterior to head, but otherwise body naked. Oesophagus cylindrical, nerve ring 172 jun from anterior. Tail cylindrical for most of its length, 5.8 cloacal diameters long.

Male: Spicules paired, equal, 1.1 cloacal diam eters long, roundly cephalate at the proxim al end and with rather indistinct ventral alae. Gubernaculum rod-shaped, 14 ^m long. Six small papillose pre-cloacal supplem ents more or less equally spaced, the anteriorm ost 147 y.m in front of the cloaca.

Discussion

The only other species with cephalic setae longer than one head diam eter are R. coronata Gerlach, 1952 and R. illgi, W ieser, 1959. In R. coronata both sets of cephalic setae are of the same length. R. illgi is described from females only, but is much larger than /?. imer in all dimensions, the buccal arm ature is more highly developed and the tail is only 3.3 anal diameters long.

TRILEPTIUM PARISETUM sp. nov.(Fig. 13)

Material studied

Holotype: 1 S , B.M. (N.H.) Reg. No. 1972: 347.

Body formulae and ratios

d- 7J t ^ 3’7*5 3,880¡Am; a = 99.5; b = 5.4; c = 23.5;ÀD ö l üt) 41

Spicule = 35 [im

Description

Cuticle w ith fain t transverse striation. Mouth surrounded by three relatively low lips each bearing a pair of stout conical labial setae 7 jxm long. Six long and four short cephalic setae situated near the posterior end of the cephalic capsule, 53 and 15 $im long. The cephalic capsule is lightly punctated round the bases of these setae. Six pairs of subcephalic setae ju st posterior to capsule, 35 + 17 am long. Ten pairs of cervical setae more posteriorly, 36 + 12 \im long (one long and one short in each pair). Short setae (9-19 [im) scattered sparsely down oesophagus length; remainder of body virtually naked. Mandibles and onchia set well forward in buccal cavity. All three of about equal size. The mandibles are quite small w ith a solid appea-

156 R.M. WARWICK AND H.M. P L A T T

rance, whilst the onchia are relatively large. Oesophagus cylindrical, nerve ring 163 (im from anterior. Tail 4.0 cloacal diam eters long, an te­rior half conical and posterior half cylindrical.

Male: Spicules paired, equal, slender and non-cephalate, only 0.85 cloacal diameters long. Gubernaculum 14 ^m long, in two halves, each term inating distally in a small plate bearing two lateral projec­tions. Pre-cloacal supplement single, cuticularised, tubular, 12 [xm long and situated 49 y.m in front of the cloaca.

\ \¡IÌ

F ig . 13Trileptium parisetum sp. nov.

A: male head; B; male tail; C: lateral view of left spicule and gubernaculum;D : lateral view of right spicule and gubernaculum.

Discussion

This species is closest to T. longisetosum Inglis, 1966. It differs from this species principally in the arrangem ent of setae on the head, the form of the gubernaculum and shape of the tail.

Acknowledgement

We would like to thank Mr. A.D. Me Intyre for providing laboratory space and equipment at Loch Ewe during our visit.

MARINE NEMATODES 157

Summary

Thirteen species of freeliving marine nematodes from the sandy beach at Firemore Bay, W ester-Ross, Scotland, are described. Amongst these we have pro­posed one new genus, Gairleanema, w ith the type species G. anagremilae gen. et sp. nov. Eleven others are new species referable to existing genera: Odontophora exharena sp. nov., Theristus (Daptonema) gelana sp. nov., Cobbia Caledonia sp. nov., Gonionchus inaequalis sp. nov., Xyala smo sp. nov., Siphonolaimus ewensis sp. nov., Metachromadora (Bradglaimus) scotlandica sp. nov., Microlaimus acinaces sp. nov., Dasynemella albaensis sp. nov., Rhabdodemania imer sp. nov., and Triliptium pari- setum sp. nov. One species (Trefusia zostericola Allgén, 1933) is redescribed, since it was hitherto poorly known.

Zusammenfassung

Dreizehn Arten von freilebenden Meeres-Nematoden, von dem sandigen Strand in Firemore Bay, W ester-Ross, Schottland, sind hier beschrieben. Unter diesen befindet sich eine neue Gattung, Gairleanema, die wir vorgeschlagen haben, m it der Type G. anagremilae gen. et sp. nov. Elf andere Arten sind neu, beziehen sich aber auf existierende genera: Odontophora exharena sp. nov., Theristus (Daptonema) gelana sp. nov., Cobbia Caledonia sp. nov., Gonionchus inaequalis sp. nov., Xyala smo sp. nov., Siphonolaimus ewensis sp. nov., Metachromadora (Bradglaimus) scotlandica sp. nov., Microlaimus acinaces sp. nov., Dasynemella albaensis sp. nov., Rhabdodemania imer sp. nov. und Trileptium parisetum sp. nov. Eine Art (Trefusia zostericola Allgén, 1933) ist wieder neu beschrieben, da dieselbe bisher nur wenig bekannt war.

REFERENCES

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C o b b , N .A ., 1920. — One hundred new nemas. (Type species o f 100 new genera).Contr. Sci. Nemat., 9, pp. 217-343.

F iL T P J E V , I .N ., 1927. — Les Nematodes libres des mers septentrionales appartenant à la fam ille des Enoplidae. Arch. Naturgesch., 91A, pp. 1-216.

G E R L A C H , s .a . , 1950. — Die Nematoden-Gattung Microlaimus. Zool. Jb. Syst., 79,pp. 188-208.

G E R L A C H , s .a ., 1952. — Nematoden aus dem Küstengrundwasser. Abh. math.-naturw.Kl. Akad. Wiss. Mainz., Jg. 1952, No. 6, pp. 1-58.

G E R L A C H , s .a . , 1953. — Die Nematodenbesiedlung des Sandstrandes und des Küsten­grundwassers an der italienischen Küste. Arch. Zool. ital., 37, pp. 519-640.

g e r l a c h , s .a . , 1955. — Zur Kenntnis der freilebenden marinen Nematoden von San Salvador. Z. wiss. Zool., 158, pp. 249-303.

g e r l a c h , s .a . , 1956. — Diagnosen neuer Nematoden aus dem Kieler Bucht. Kieler Meeresforsch., 12, pp. 85-109.

h o p p e r , B .E ., 1968. — Marine nematodes of Canada. I. Prince Edward Island. Can.J. Zool., 46, pp. 1103-1111.

i N G L i s , W .G ., 1966. — Marine nematodes from Durban, South Africa. Bull. Br. Mus.nat. Hist. {Zool.), 14, pp. 79-106.

d e m a n , J .G . , 1907. — Sur quelques espèces nouvelles ou peu connues de Nématodes libres habitant les côtes de la Zélande. Mêm. Soc. Zool. France, 20, pp. 33-90.

M c I n t y r e , a .D . and E L E F T H E R io u , A ., 1968. — The bottom fauna of a fla tfish nursery ground. / . mar. biol. Ass. U.K., 48, pp. 113-142.

R iE M A N , f . , 1966. — Die interstitielle Fauna im Elbe-Aestuar. Verbreitung und Sys­tem atik. Arch . Hydrobiol., Suppl. 31, pp. 1-279.

REM A N N , f . , 1966a. — Die Gattung Trefusia de Man, 1893 (Enoplida : Oxystom inidae). Veröff. Inst. Meeresforsch. Bremerh., 10, pp. 1-29.

158 i?.ili. WARWICK AND H M . P L A T T

Vi t i e l l o , p . , 1969. — Linhomoeidae (Nematoda) des vases profondes du Golfe du Lion. Téthys, 1, pp. 493-527.

w i e s e r , w., 1953. — Freeliving marine nematodes I. Enoploidea. Reports of the Lund University Chile Expedition 1948-49. Acta Univ. Lund. N.F., 49 (6), pp. 1-155.

w i e s e r , w., 1956. — Freeliving marine nematodes III. Axonolaim oidea and Mon- hysteroidea. Reports of the Lund University Chile Expedition 1948-49. Acta Univ. lund, N.F., 53(13), pp. 1-115.

w i e s e r , w., 1959. — Freeliving marine nematodes and other sm all invertebrates of Puget Sound beaches. Univ. Wash. Pubis Biol ., 19, pp. 1-179.

w i e s e r , w. and h o p p e r , b .e ., 1967. — Marine nematodes of the East Coast of North America I. Florida. Bull. Mus. comp. Zool. Harv ., 135, pp. 239-344.

Note added in proof

During the period in which this paper was in press, Ward (1972) has described two species of Xyala from Liverpool Bay. His X. longicaudata is clearly identical w ith X. smo and is thus the preferred name, X. smo becoming a nomen nudum.

REFERENCE: w a r d , a . r . , 1972. — Two new species of Xyala (Nematoda, Monhysteroidea) from sublittoral sediments in Liverpool Bay. Mar. Biol., 13 (2), pp. 176-178.